Positive-single stranded RNA viruses comprising the Retroviridae family include those of the subfamily Orthoretrovirinae and genera Alpharetrovirus, Betaretrovirus, Gamaretrovirus, Deltaretrovirus, Epsilonretrovirus, Lentivirus, and Spumavirus which cause many human and animal diseases. Among the Lentivirus, HIV-1 infection in humans leads to depletion of T helper cells and immune dysfunction, producing immunodeficiency and vulnerability to opportunistic infections. Though progress has been made in treating HIV-1 infections (Hammer, S. M., et al.; JAMA 2008, 300: 555-570), HIV infections remain a global health concern. As such, there remains a pressing need to discover new antiretroviral agents that are active against HIV.